SELECTIVE FORAGING BY BEAVERS FOR TREES SIZE AND DISTANCE FROM WATER Justin Bochnak, Mammalogy EEOB 625,

, The Ohio State University, 27 February, 2012

Introduction The foraging behavior of the American beaver (Castor canadensis) has caused the species to be the subject of a significant amount of research. Much of this research examines how beavers change the structure and function of the areas they occupy. Beavers are considered ecosystem engineers because their foraging behavior can result in modifications that transform the adjoining ecosystem (Voelker and Dooley, 2008). Determining if beavers show a selective preference for certain species or sizes of trees is important to understand what transformations, within local ecosystems, are likely to result from beaver foraging activity (Voelker and Dooley, 2008). Each species of woody plant has different structural and nutritional properties that are likely to affect the decisions of selectively foraging beavers. Studying beaver foraging behavior not only increases the scientific understanding of how beavers impact the ecosystem of the areas they occupy, but it also makes the species ideal for examining the optimal foraging and central place foraging theories. The optimal foraging theory predicts that consumers become progressively more selective, for higher quality resources, as the distance required to obtain these resources increases (Voelker and Dooley, 2008). Central place foraging theory applies the assumptions of an optimal foraging individual to species that transport harvested resources back to a central location (Voelker and Dooley, 2008). A beavers diet consists explicitly of herbaceous material, a majority of which is harvested on land and transported back to their den for consumption

(Fryexll and Doucet, 1990; Raffel et al., 2009). This makes the central place foraging theory particularly relevant for studying a species such as the American beaver. By examining which trees beavers cut as they travel further from shore, researchers are able to test the central foraging theory gaining insight that has that has improved the scientific understanding of community and trophic interactions for many different species (Hyonen and Nummi, 2008). To better understand the implications of beaver foraging behavior, the objectives of this study are to evaluate if the diameter of trees harvested change with the distance traveled from shore, and to determine if there is a selective preference, by beavers, for specific tree species within the forest plant community. I predict that tree diameter of harvested trees will increase with increasing distance traveled from shore, and that beavers will selectively harvest a disproportionate amount of certain tree species within the woody plant community.

Methods Studies were conducted during January and February, 2012 at two locations; Mudhen Marsh in Sunbury, Ohio and Delaware State Park in Delaware, Ohio. At each of the study locations the sample plots were placed at a distance of ten and thirty meters from shore; designating near and far plots, respectively. A total of six plots, three near and three far, were sampled at the Delaware State Park location. At Mudhen Marsh, measurements were recorded in four near and four far plots for a total of eight sample plots. Each plot measured ten by ten meters and was positioned at a distance of ten meters from adjacent sample plots. Within each plot, measurements were recorded for the number of trees, the number of cut and uncut trees, tree diameter, and tree species. Trees were considered cut only if they 2

were completely cut, otherwise they were counted as uncut trees. Tree diameter measurements were taken between 15 and 30 cm above ground depending on the condition of the tree and the height at which it was cut. Species was recorded for each tree in the sample plots, and then categorized into one of four groups; Acer species, Cratageous species, Ulmus species, and other species. The data was analyzed to determine if the beavers showed a selective preference for specific species or sizes of trees. A t-test was used to evaluate differences in mean diameter of cut and uncut trees located in the near and far plots. Chi-square tests were utilized to assess the relative proportion of cut and uncut trees in the near and far plots for each of the species categories.

Results The relative proportions of trees available (cut: uncut: total) were similar between the two sample sites (n Mudhen = 30: 106: 136, n Deleware = 30: 66: 96) (Fig. 1). Both sites had a greater number of trees in the near sample plots (n Mudhen =78, n Delaware = 68) relative to the far plot locations (n Mudhen = 58, n Delaware = 28) (Fig. 2). At Mudhen Marsh, of the total trees, 78 percent were uncut. At Delaware State Park, of the total trees, 69 percent were uncut. At Mudhen Marsh the forest community was composed primarily of Ulmus and Carya species. At Delaware State Park the forest community was composed mainly of Acer, Carya, and Ulmus species. Beavers preferentially cut some tree species compared to others at both study sites (Mudhen X2= 8.9, p < 0.02; Delaware X2 = 8.5, p < 0.02). At both study sites, trees categorized as 3

other composed the greatest percentage of the forest community, but contained the smallest fraction of cut trees (Fig. 3 and 4). At Delaware, beaver did not cut all tree species in equal proportions (X2=8.5, p <0.02), but selectively cut trees within the Acerace group (Fig. 4). This indicates that the beavers, at Delaware State Park, may be exhibiting a selective preference for certain tree species. Beaver at Mudhen Marsh cut trees that were significantly larger in the far plots compared to the near (t=7.74, p < 0.001) (Fig. 5). This trend was not observed from the data collected at Delaware State Park; instead, trees of a similar, intermediate, diameter were harvest from both near and far plots (t=0.223, p=0.85) (Fig. 6).

Discussion At the Mudhen Marsh study site, data analysis determined that the beavers at this location displayed a size selective preference for larger diameter trees as they traveled further from shore. The significantly larger mean diameter of cut trees in the far plots indicates that the beavers at this location were foraging for higher quality resources, consistent with the predictions of the optimal and central place foraging theories. However, the data from Delaware State Park did not show the same size selective patterns. Instead, the data recorded at Delaware State Park suggested that the beavers were foraging on intermediate sized trees in both the near and far plots. Similar mean diameter for cut trees in the near and far plots implies an absence of selective preference for higher quality resources by beavers foraging at this location. Such data may have resulted from differences in the physical structure or species

composition of the forest plant community at this location. Such foraging behavior cannot be considered consistent with the predictions of either foraging theories. The conflicting results derived at during this study characterize much of the existing research examining beaver foraging behavior. Several studies have confirmed size selective foraging, consistent with the predictions outlined by the optimal foraging theory (Fryxell and Doucet, 1993; Vokelker and Dooley, 2008; Raffel et al., 2009). However, a significant amount of research has also failed to recognize patterns of such behavior (Jenkins, 1980; Fryxell and Doucet, 1993; Hyvonen and Nummi, 2008). The cause for inconsistent results associated with the research related to beaver foraging behavior has been a subject of significant speculation. The differential nutritional quality associated with distinctive tree species has been suggested as one reason why beavers might choose certain trees over others despite size or distance from shore (Fryxell and Doucet, 1990). The increased energetic expense of transporting larger trees over longer distances has also been identified as a possible explanation for why beavers might cut smaller trees as they travel further from shore (Jenkins, 1980; Hyvonen and Nummi, 2008). Several hypothetical justifications have proven reasonable. However, science suggests a combination of factors resulting from differences in forest composition and resource abundance within distinct systems determines the selective foraging behavior of beavers (Fryxell and Doucet, 1993).

Acknowledgements I would like to thank my classmates Todd Karg and Sam Carsile who helped in the data collection and configuration information and figures presented in this paper. My TA Sarah Smiley, who helped explain the data collection and analysis processes, and Dr. John Harder, the instructor of the course.

Literature Cited Barnes, D. M., and A. U. Mallik. 2001. Effects of Beaver, Castor canadensis, herbivory on streamside vegetation in a northern Ontario watershed, Canadian Field-Naturalist 115:9-21. Fryxell, J. 1999. Functional responses to resource complexity: an experimental analysis of foraging by beavers. Fryxell, J. M., and C. M. Doucet. 1991. Provisioning time and central-place foraging in beavers, Canadian Journal of Zoology-Revue Canadienne De Zoologie 69:1308-1313. Fryxell, J. M., and C. M. Doucet. 1993. Diet choice and functional response curves, Ecology 74:1297-1306. Hyvonen, T., and P. Nummi. 2008. Habitat dynamics of beaver Castor canadensis at two spatial scales, Wildlife Biology 14:302-308. Raffel, T. R., N. Smith, C. Cortright, and A. J. Gatz. 2009. Central Place Foraging by Beavers (Castor canadensis) in a Complex Lake Habitat, American Midland Naturalist 162:62-73. Voelker, B. W., and J. L. Dooley, JR. 2008. Impact by North American beaver (Castor canadensis) on forest plant composition in the wilds, a surface-mined landscape in southeastern Ohio, Ohio Journal of Science 108:9-15.

Tables and Figures

160 140 120 Number of Trees 100 80 60 40 20 0 Mudhen Marsh Site Location Delaware State Park Cut Uncut Total

Figure 1. The trees available at the Mudhen Marsh and Delaware State Park sample sites. The proportion of total trees, number of trees cut, and number of trees uncut are represented for each study site. At Mudhen Marsh data was collected from a total of eight plots; four near and four far. Delaware data was collected over six total plots; three near and three far.

40 Average Diameter (cm.) 35 30 25 20 15 10 5 0

n = 68 n = 29 n = 78 n = 58

Near Far

Mudhen Marsh Site Location

Delaware State Park

Figure 2. Average tree diameter for all trees sampled in the near and far plots at Mudhen Marsh and Delaware State Park. At both locations the average diameter of trees in the near plots exceeded the average diameter of trees in the far plots. Error bars represent the standard deviation. 7

40 Cut 35 30 Number of Trees 25 20 15 10 5 0 Acer spp. Carya/Ulmus spp. Tree species other Uncut

Figure 3. The number of cut and uncut trees located in the near and far plots for each of the different groups of species examined at Delaware State Park.

60 50 Number of Trees 40 30 20 10 0 Ulmus spp. Carya spp. Tree species other

Cut Uncut

Figure 4. The number of cut and uncut trees located in the near and far plots for each of the different groups of species examined at Mudhen Marsh.

45 40 Average tree diameter (cm.) 35 30 25 20 15 10 5

n = 22 n = 56 n=8 n = 50

Cut Uncut

0 Near Mudhen Marsh Far

Figure 5. Mean diameter of cut and uncut trees located in the near and far plots at Mudhen Marsh. The error bars represent the standard error of the mean.
40 Cut 35 Average tree diameter (cm.) 30 25 20 15 10 5
n = 23 n = 45 n=7 n = 21

Uncut

0 Near Delaware State Park Far

Figure 6. Average tree diameter of cut and uncut trees in the near and far plots at Delaware State Park. Error bars represent standard error of the mean.